发育与进化
（Evolutionary developmental biology，常简称Evo-Devo）
王 文
中德Max Planck青年科学家小组
遗传资源与进化国家重点实验室
中国科学院昆明动物研究所
内容：
1、进化发育生物学简介
2、我们的思考和研究
做科学要有好的科学问题：
科学体系发展中不可回避、必须回答的问题
可以操作的问题
The Art of feasible!
背景：
• 遗传、发育和进化是生命科学中的本质问题，我们看到的各种各样的
生物组织、器官和性状无不是千百万年来遗传变异经自然选择固定并
最后通过个体发育展现而来。长期以来科学家们一直在试图把这三方
面联系起来。随着分子生物学、进化基因组学和发育生物学等现代生
命科学和技术的迅速发展和结合，这种努力首次变得富有希望。20多
年前兴起的进化发育生物学（Evolutionary developmental biology
，常简称Evo-Devo）是这种希望的一个开端，该学科试图从基因表达
在发育过程中的变化理解不同物种形态发生和功能的进化(Hall，1992
）。
Haeckel‘s theory:
Ontogeny recapitulates phylogeny （1874）
• the fetal development of an individual (ontogeny) is a speeded-up replay
of millions of years of species evolution (phylogeny)
Thomas Hunt Morgan, “Embryology and Genetics” (1934)
第一次提出了胚胎发育的可能细胞分化机制：基因表达的时空分
化
在过去的30年中，我们已经发现了各种动物均使用的发育调控基
因基本的细胞过程，形成了“进化发育生物学”这一分支学科。
进化发育生物学（evolutionary developmental biology）：
旨在阐明生物进化中发育过程和机制怎样改变以及这些改变如何造就过去
和现在的生物多样性的一门学科。（Baguna and Garcia-Fernandez, 2003).
物种之间发育基因的比较（Sean Carrol， 2005）
常简称“Evo Devo”
"Evo-devo" is one of the newest areas of biology. It seeks to explain
evolutionary events through the mechanisms of developmental biology. Its
major questions are often the questions of the late 1800s which had been left
waiting for other generations of biologists to unravel. We are those
generations. These questions include: How did wings evolve? How did
limbs evolve? How did vertebrates arise from invertebrates? How did
vertebrates form their head?
The Modern Synthesis of the 1930s and 1940s was the synthesis of
evolutionary biology and population genetics. It is one of the greatest
explanatory systems in science, and it explains incredibly well the origin,
maintenance, and evolution of variations within populations of species.
However, it did not identify the genes involved in these changes, nor did it
explain the origin of higher taxa. Evolutionary developmental biology seeks
to “complete” the synthesis by adding developmental genetics to population
genetics. Traditional Evo-devo looks at the regulatory regions of genes
rather than their coding sequences. It emphasizes the arrival of the fittest
rather than the survival of the fittest.
群体遗传学案例（基因在群体中的变异情况）
Polymorphism of gene toy and sphinx in a worldwide and a local Drosophila
melanogaster population
_____________________________________________________________________________________________________________________________________________________________
Stocks
Locations
toy intron1(847bp)
locus sphinx (4257bp)
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1 1 1111111111111 1 1 1 1 1 1 2 2 2 2 2 2 2 3 33 3 3 3 4
Position
1 1 1 1 1 2 3 4 4 5 5 7
1 1 11111111122 2 2 5 5 5 555555555555556666 6 8 9 1 2 7777777777777 7 7 7 8 9 9 0 1 2 2 2 5 7 0 11 1 8 9 0
1 4 8 8 9 0 4 4 6 1 3 5
9 1 3 99999999900 1 7 1 1 2 888899999999990000 5 1 2 3 9 2223333333334 5 8 9 8 1 3 3 4 2 4 4 4 0 0 33 7 0 1 0
3 1 0 1 3 4 2 2 2 0 7 8
0 6 0 12345678901 0 6 5 8 2 678901234567890123 4 5 0 8 9 7890124567890 5 1 9 7 9 7 4 7 6 1 5 0 6 8 67 1 7 5 4
closs19
closs20
253.4
253.27
Y2
Y10
Yep25
south1.10
southA.9
northN34
prunay
Rio
ok17
Hg84
LA79
ZH56
ZW30
ZW56
closs16
north7.13
253.38
253.35
253.30
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World
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ATTTTACAAAA
----------ATTTTACAAAA
population
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New York popalation (Upstate of New York, USA)
closs3
USA
A A A C C T G C T A C C
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closs4
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closs7
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closs20
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closs27
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closs28
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______________________________________________ ______________________________________________________________________________________________________________
Wang, W., Thornton, K., Berry, A. and Long, M. 2002, Science, 295: 134-137.
No crossing over
Subject to relentless selective sweep and background selection
in populations
Drosophila
Chromosome
Variations in 18 gene regions were investigated
Wang, W., Thornton, K., Berry, A. and Long, M. 2002, Science, 295: 134-137.
Evo-Devo 中的重要概念:
(1) Phylogeny (系统发育)
Evo-Devo 中的重要概念:
(2) Body plan
胚胎前后轴（A－P）、背腹轴（D－V）
、左右轴（L－R）等决定
De Robertis, 2008)
Body
plan
Evo-Devo 中的重要概念:
(3) Toolkit genes: master genes that govern the formation and patterning of
their
Bodies and body parts of all kinds of animals.
Despite being morphologically very diverse, multicellular organisms
are made by a very conserved set of regulatory genes (tool kit genes) playing
comparable developmental roles. This most unexpected finding
represented a powerful molecular proof of evolution as ‘descent
with modification’ (Darwin, 1859).
最近：
A set of genes that is required for development and is shared widely among
species (Canestro et al, NRG, 2007)
传统的发育进化生物学研究范式：经典的进化发育生物学 (Evo
Devo) 认为所有动物的发育由同一套工具箱基因 （toolkit
genes） 控制
在果蝇，ubx抑制腹部dsl基因的表达的足产生，但在蜈蚣中其
的C端磷酸化位点不存在，导致不抑制腹部的足产生
果蝇
蜈蚣
不同动物眼睛的发育控制由一套同源的基因（Pax6）控制
Nonetheless, it brings a big paradox: if
developmental genes are the same, how
are differences in development and in the
final morphology in different organisms
to be accounted for?
(Canesttro et al., NRG, 2007)
(Canesttro et al., NRG, 2007)
最近，Sean
Carroll等认为
发育进化主要
由调控序列的
改变导致
Loss of an Enhancer in Stickleback Fish Evolution
leads to dramatic morphological difference
Butterfly Wing Pattern Evolution Is Associated with Changes in a
Notch/Distal-less Temporal Pattern
Current biology 2004, 13:1159-1166
(4) Homology vs. convergent, parallel and homoplasmy
Sorting out parallelism from homology in landmark issues such as the origin(s)
of segmentation, the origin(s) of the eye (Hodin, 2000) warrants
further studies of additional phyla, and a deep understanding of
plasticity and constraints in gene and developmental evolution.
When a structure which is homologous between
closely related organisms is built using different genes, it should be
considered convergent rather than homologous.
Homologous genes may be responsible for non-homologous
morphologies. Genes and pathways are utilized on many separate
occasions during development.
As examples, the Notch-signalling pathway is broadly used
during Drosophila and vertebrate development (Simpson, 1997;
Robey, 1997), and hedgehog, TGF-β and Wnt family members are
used over and over again during development. Evolution may well
have worked by “genetic tinkering” (Jacob, 1977) or “bricolage” of
gene networks (Duboule and Wilkins, 1998).
paralogues, orthologues
Paralogues:源自基因重复；共同祖先是基因。
Orthologues：源自祖先物种，共同祖先是物种。
Macroevolution vs. microevolution
As Dobzhansky (1937) firstly pointed out, the main issue in the
macro vs. microevolution debate is whether mutations resulting in
real evolutionary novelties are of the same kind as those occurring
daily or whether we should expect special, rare mutations only
occurring on geological time scales.
Evo Devo使用的主要方法
1、比较基因组学
2、各种生物胚胎发育中基因的表达图式
3、同源基因的替换或转基因
The future task for macro-Evo-Devo will be to unravel: i) preexisting
developmental potential; in other words, what was before
the Cambrian, which means analyzing the developmental toolkit
component of the closest sister groups, relatives of eubilaterians,
the acoelomorph flatworms (Ruiz-Trillo et al., 1999; 2002); ii) the
extent and quality of bricolage of this basal bilaterian toolkit
compared to higher bilaterians; and iii) trying to link the known, and
those which remain to be discovered, fossil groups
Ongoing debates or unsolved questions in evolutionary developmental biology
Origin of Bilateria
Origin of Vertebrata
Origin of Birds
Origin of Limbs
Origin of Brain
·但传统的进化发育生物学更多是探讨动物中古老的、共有的发育机制，很多
看法和假说难以在物种分化的水平得到验证和阐释。
·近年来Sean Carroll（2005）等开始研究近缘种间形态变异的基因调控机制
，并得出了形态变异主要是基因调控方式改变的结论。
·但根据我们过去一些年来研究新基因及其新功能起源进化的结果看，上述两
个Evo-Devo的研究范式可能处在了两个极端:
一个考察太过古老的事件因而容易流于争辨；
一个过于强调小的变异(regulatory sequence )而忽略基因组中大的、
重要的遗传改变的作用。
Evo Devo
• Evolutionary development is so new
there isn’t even consensus as to
how to spell it.
• We spell it “evo devo”
because that’s how Sean
Carroll spells it.
• Others spell it “evo-devo”
and “evodevo.”
植物的发育与进化
2006年
2006
2007年
2007年